Investigating flock-associated mimicry: examining the evidence for, and drivers of, plumage mimicry in the greater and lesser necklaced laughingthrush.

Visual mimicry is less understood in birds than in other taxa. The interspecific social dominance mimicry (ISDM) hypothesis asserts that subordinate species resemble dominant ones to reduce aggression. Plumage mimicry has also been consistently noted in mixed-species flocks (MSFs), suggesting a connection to grouping behaviour, although it is unclear whether this is linked to ISDM. We studied greater necklaced laughingthrush (GNLT, Pterorhinus pectoralis) and lesser necklaced laughingthrush (LNLT, Garrulax monileger), which were recently placed in different genera. Measurements of 162 museum specimens showed LNLT converging in sympatry with GNLT in necklace colour, but diverging in necklace to body ratio, with proportionally smaller necklaces. The species were closely associated in six of seven MSF systems from Nepal to China. In a study of foraging behaviour in Nepal, aggression was rare between the species, LNLT followed GNLT and had lower foraging rates when further from GNLT. Our data suggest a link between this MSF-associated mimicry and ISDM, and that the subordinate LNLT may be the mimic and gain more from the resemblance. The species spend much time together in dense and poorly lit vegetation, where the LNLTs resemblance to GNLTs potentially allows them to forage closer to GNLTs than would be otherwise possible.

Exploring potential relationships between acoustic indices and ecosystem functions: a test on insect herbivory.

Biodiversity loss is a global concern. Current technological advances allow the development of novel tools that can monitor biodiversity remotely with minimal disturbance. One example is passive acoustic monitoring (PAM), which involves recording the soundscape of an area using autonomous recording units, and processing these data using acoustic indices, for example, to estimate the diversity of various vocal animal groups. We explored the hypothesis that data obtained through PAM could also be used to study ecosystem functions. Specifically, we investigated the potential relationship between seven commonly used acoustic indices and insect leaf herbivory, measured as total leaf damage and as the damage from three major insect feeding guilds. Herbivory was quantified on seedlings in 13 plots in four subtropical forests in south China, and acoustic data, representing insect acoustic complexity, were obtained by recording the evening soundscapes in those same locations. Herbivory levels correlated positively with the acoustic entropy index, commonly reported as one of the best-performing indices, whose high values indicate higher acoustic complexity, likely due to greater insect diversity. Relationships for specific feeding guilds were moderately stronger for chewers, indicating that the acoustic indices capture some insect groups more than others (e.g., chewers include soniferous taxa such as crickets, whereas miners are mostly silent). Our findings suggest that the use of PAM to monitor ecosystem functions deserves to be explored further, as this is a research field with unexplored potential. Well-designed targeted studies could help us better understand how to best use novel technologies to monitor ecosystem functions.

Crop and landscape heterogeneity increase biodiversity in agricultural landscapes: A global review and meta-analysis.

Agricultural intensification not only increases food production but also drives widespread biodiversity decline. Increasing landscape heterogeneity has been suggested to increase biodiversity across habitats, while increasing crop heterogeneity may support biodiversity within agroecosystems. These spatial heterogeneity effects can be partitioned into compositional (land-cover type diversity) and configurational heterogeneity (land-cover type arrangement), measured either for the crop mosaic or across the landscape for both crops and semi-natural habitats. However, studies have reported mixed responses of biodiversity to increases in these heterogeneity components across taxa and contexts. Our meta-analysis covering 6397 fields across 122 studies conducted in Asia, Europe, North and South America reveals consistently positive effects of crop and landscape heterogeneity, as well as compositional and configurational heterogeneity for plant, invertebrate, vertebrate, pollinator and predator biodiversity. Vertebrates and plants benefit more from landscape heterogeneity, while invertebrates derive similar benefits from both crop and landscape heterogeneity. Pollinators benefit more from configurational heterogeneity, but predators favour compositional heterogeneity. These positive effects are consistent for invertebrates and vertebrates in both tropical/subtropical and temperate agroecosystems, and in annual and perennial cropping systems, and at small to large spatial scales. Our results suggest that promoting increased landscape heterogeneity by diversifying crops and semi-natural habitats, as suggested in the current UN Decade on Ecosystem Restoration, is key for restoring biodiversity in agricultural landscapes.

High association strengths are linked to phenotypic similarity, including plumage color and patterns, of participants in mixed-species bird flocks of southwestern China.

Participants in mixed-species bird flocks (MSFs) have been shown to associate with species that are similar in body size, diet, and evolutionary history, suggesting that facilitation structures these assemblages. In addition, several studies have suggested that species in MSFs resemble each other in their plumage, but this question has not been systematically investigated for any MSF system. During the nonbreeding season of 2020 and 2021, we sampled 585 MSFs on 14 transects in 2 habitats of Tongbiguang Nature Reserve in western Yunnan Province, China. We performed social network analysis and the Multiple Regression Quadratic Assignment Procedure to evaluate the effect of 4 species traits (body size, overall plumage color, distinctive plumage patterns, and diet) and evolutionary history on species association strength at the whole-MSF and within-MSF levels. All 41 significant relationships showed that species with stronger associations were more similar in their various traits. Body size had the strongest effect on association strength, followed by phylogeny, plumage patterns, and plumage color; diet had the weakest effect. Our results are consistent with the hypotheses that the benefits of associating with phenotypically similar species outweigh the potential costs of interspecific competition, and that trait matching can occur in plumage characteristics, albeit more weakly than in other traits. Several explanations exist as to why similarities in plumage may occur in MSFs, including that they could reduce predators' ability to target phenotypically "odd" individuals. Whether trait matching in plumage occurs through assortative processes in ecological time or is influenced by co-evolution requires further study.

Species diversity and interspecific information flow.

Interspecific information flow is known to affect individual fitness, population dynamics and community assembly, but there has been less study of how species diversity affects information flow and thereby ecosystem functioning and services. We address this question by first examining differences among species in the sensitivity, accuracy, transmissibility, detectability and value of the cues and signals they produce, and in how they receive, store and use information derived from heterospecifics. We then review how interspecific information flow occurs in communities, involving a diversity of species and sensory modes, and how this flow can affect ecosystem-level functions, such as decomposition, seed dispersal or algae removal on coral reefs. We highlight evidence that some keystone species are particularly critical as a source of information used by eavesdroppers, and so have a disproportionate effect on information flow. Such keystone species include community informants producing signals, particularly about predation risk, that influence other species' landscapes of fear, and aggregation initiators creating cues or signals about resources. We suggest that the presence of keystone species means that there will likely be a positive relationship in many communities between species diversity and information through a 'sampling effect', in which larger pools of species are more likely to include the keystone species by chance. We then consider whether the number and relative abundance of species, irrespective of the presence of keystone species, matter to interspecific information flow; on this issue, the theory is less developed, and the evidence scant and indirect. Higher diversity could increase the quantity or quality of information that is used by eavesdroppers because redundancy increases the reliability of information or because the species provide complementary information. Alternatively, there could be a lack of a relationship between species diversity and information if there is widespread information parasitism where users are not sources, or if information sourced from heterospecifics is of lower value than that gained personally or sourced from conspecifics. Recent research suggests that species diversity does have information-modulated community and ecosystem consequences, especially in birds, such as the diversity of species at feeders increasing resource exploitation, or the number of imitated species increasing responses to vocal mimics. A first step for future research includes comprehensive observations of information flow among different taxa and habitats. Then studies should investigate whether species diversity influences the cumulative quality or quantity of information at the community level, and consequently ecosystem-level processes. An applied objective is to conserve species in part for their value as sources of information for other species, including for humans.

Examining the co-occurrences of human threats within terrestrial protected areas.

Human threats to biodiversity are prevalent within protected areas (PAs), undermining their effectiveness in halting biodiversity loss. Certain threats tend to co-occur, resulting in amplified cumulative impact through synergistic effects. However, it remains unclear which threats are related the most. We analyzed a dataset of 71 human threats in 18 013 terrestrial PAs of the European Union's Natura 2000 network, using a Joint Species Distribution Modelling approach, to assess the threats' co-occurrence patterns and potential drivers. Overall, threats were more frequently correlated positively than negatively. Threats related to agriculture and urbanization were correlated strongly with most other threats. Approximately 70% of the variance in our model was explained by country-specific factors, indicating the importance of local drivers. Minimizing the negative impact of key threats can likely reduce the impact of related threats. However, more research is needed to understand better the relationships among threats and, importantly, their combined impact on biodiversity.

Ecological filtering shapes the impacts of agricultural deforestation on biodiversity.

The biodiversity impacts of agricultural deforestation vary widely across regions. Previous efforts to explain this variation have focused exclusively on the landscape features and management regimes of agricultural systems, neglecting the potentially critical role of ecological filtering in shaping deforestation tolerance of extant species assemblages at large geographical scales via selection for functional traits. Here we provide a large-scale test of this role using a global database of species abundance ratios between matched agricultural and native forest sites that comprises 71 avian assemblages reported in 44 primary studies, and a companion database of 10 functional traits for all 2,647 species involved. Using meta-analytic, phylogenetic and multivariate methods, we show that beyond agricultural features, filtering by the extent of natural environmental variability and the severity of historical anthropogenic deforestation shapes the varying deforestation impacts across species assemblages. For assemblages under greater environmental variability-proxied by drier and more seasonal climates under a greater disturbance regime-and longer deforestation histories, filtering has attenuated the negative impacts of current deforestation by selecting for functional traits linked to stronger deforestation tolerance. Our study provides a previously largely missing piece of knowledge in understanding and managing the biodiversity consequences of deforestation by agricultural deforestation.

A multidimensional framework to quantify the effects of urbanization on avian breeding fitness.

Urbanization has dramatically altered Earth's landscapes and changed a multitude of environmental factors. This has resulted in intense land-use change, and adverse consequences such as the urban heat island effect (UHI), noise pollution, and artificial light at night (ALAN). However, there is a lack of research on the combined effects of these environmental factors on life-history traits and fitness, and on how these interactions shape food resources and drive patterns of species persistence. Here, we systematically reviewed the literature and created a comprehensive framework of the mechanistic pathways by which urbanization affects fitness and thus favors certain species. We found that urbanization-induced changes in urban vegetation, habitat quality, spring temperature, resource availability, acoustic environment, nighttime light, and species behaviors (e.g., laying, foraging, and communicating) influence breeding choices, optimal time windows that reduce phenological mismatch, and breeding success. Insectivorous and omnivorous species that are especially sensitive to temperature often experience advanced laying behaviors and smaller clutch sizes in urban areas. By contrast, some granivorous and omnivorous species experience little difference in clutch size and number of fledglings because urban areas make it easier to access anthropogenic food resources and to avoid predation. Furthermore, the interactive effect of land-use change and UHI on species could be synergistic in locations where habitat loss and fragmentation are greatest and when extreme-hot weather events take place in urban areas. However, in some instances, UHI may mitigate the impact of land-use changes at local scales and provide suitable breeding conditions by shifting the environment to be more favorable for species' thermal limits and by extending the time window in which food resources are available in urban areas. As a result, we determined five broad directions for further research to highlight that urbanization provides a great opportunity to study environmental filtering processes and population dynamics.

Interspecific sociality alters the colonization and extinction rates of birds on subtropical reservoir islands.

Island biogeography theory has proved a robust approach to predicting island biodiversity on the assumption of species equivalency. However, species differ in their grouping behaviour and are entangled by complex interactions in island communities, such as competition and mutualism. We here investigated whether intra- and/or interspecific sociality may influence biogeographic patterns, by affecting movement between islands or persistence on them. We classified bird species in a subtropical reservoir island system into subcommunities based on their propensity to join monospecific and mixed-species flocks. We found that subcommunities which had high propensity to flock interspecifically had higher colonization rates and lower extinction rates over a 10-year period. Intraspecific sociality increased colonization in the same analysis. A phylogenetically corrected analysis confirmed the importance of interspecific sociality, but not intraspecific sociality. Group-living could enable higher risk crossings, with greater vigilance also linked to higher foraging efficiency, enabling colonization or long-term persistence on islands. Further, if group members are other species, competition can be minimized. Future studies should investigate different kinds of island systems, considering positive species interactions driven by social behaviour as potential drivers of community assembly on islands. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

Can convergence in mixed-species flocks lead to evolutionary divergence? Evidence for and methods to test this hypothesis.

One of the most fundamental goals of modern biology is to achieve a deep understanding of the origin and maintenance of biodiversity. It has been observed that in some mixed-species animal societies, there appears to be a drive towards some degree of phenotypic trait matching, such as similar coloration or patterning. Here we build on these observations and hypothesize that selection in mixed-species animal societies, such as mixed-species bird flocks, may drive diversification, potentially leading to speciation. We review evidence for possible convergent evolution and even outright mimicry in flocks from southwestern China, where we have observed several cases in which species and subspecies differ from their closest relatives in traits that match particular flock types. However, understanding whether this is phenotypic matching driven by convergence, and whether this divergence has promoted biodiversity, requires testing multiple facets of this hypothesis. We propose a series of steps that can be used to tease apart alternative hypotheses to build our understanding of the potential role of convergence in diversification in participants of mixed-species societies. Even if our social convergence/divergence hypothesis is not supported, the testing at each step should help highlight alternative processes that may affect mixed-species flocks, trait evolution and possible convergence. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes.

Mixed-species groups of birds, fishes and mammals have traditionally been described in taxa-specific journals. However, mixed-species systems are actually more widely found when one includes aggregative (non-moving) systems, such as those common in amphibians and invertebrates. The objective of this special issue is to dispel the idea that mixed-species phenomena are a 'niche topic' to ecology and instead explore how taking a mixed-species perspective can change our conception of important ecological patterns and processes. A mixed-species perspective starts by understanding the relative abundance and positioning of individuals of different species and their behavioural synchrony; it is enriched by understanding differences between species in their vulnerability/attractiveness to predators, their potential for competing with other group participants and their use as a source of public information. Contributions to the special issue show how the mixed-species perspective can change our ideas about invasion ecology, island biogeography, keystone species, mimicry, predator eavesdropping and more. Rather than seeking synthesis, the special issue celebrates the taxonomic and conceptual breadth of the field of mixed-species groups, with detailed descriptions of many distinctive systems. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

Audience effects in a group-living bird: How contact call rate is affected by vegetation and group size and composition.

Contact calling is a ubiquitous behavior of group-living animals. Yet in birds, beyond a general connection with group cohesion, its precise function is not well-understood, nor is it clear what stimulates changes in contact call rate. In an aviary experiment, we asked whether Swinhoe's White-eyes, Zosterops simplex, would regulate their own production of contact calls to maintain a specific rate at the group level. Specifically, we hypothesized that the sudden cessation of the group-level call rate could indicate an immediate predation threat, and we predicted that birds in smaller groups would call more to maintain a high call rate. We also investigated the effects of environmental characteristics, such as vegetation density, and social stimuli, such as the presence of certain individuals, on the rate of three different contact call types. To calculate mean individual-level rates, we measured the group-level rate and divided it by the number of birds in the aviary. We found that the individual-level rate of the most common call types increased with a greater group size, the opposite pattern to what would be expected if birds were maintaining a specific group-level rate. Vegetation density did not affect any call rate. However, individual-level rates of all call types decreased when birds were in subgroups with individuals of differing dominance status, and the rate of some call types increased when birds were with affiliated individuals. Our results do not support the hypothesis that contact calls are related to habitat structure or immediate predation risk. Rather, they appear to have a social function, used for communication within or between groups depending on the call type. Increases in call rates could recruit affiliated individuals, whereas subordinates could withhold calls so that dominants are unable to locate them, leading to fluctuations in contact calling in different social contexts.

Metal and metalloid sources apportionment in soil of two major agroecosystems of southern China.

Apportioning the sources of metals/metalloids is a critical step toward soil quality protection and ecological restoration. The objective of this study was to identify the potential sources of contamination of As, Cd, Cr, Cu, Hg, Mn, Pb, and Zn, and determine the contribution rates of each source, to rice and sugarcane agroecosystems of southwestern Guangxi, southern China. We collected a total of 300 soil samples at a former lead-zinc mine and at two reference sites, 6 and 60 km away from the mine, sampling both agroecosystems at each site. Overall, the positive matrix factorization (PMF) receptor model revealed that in rice paddies at the mine site, mining activities had the highest contribution (60.7% of all examined metals/metalloids), followed by irrigation (25.8%), and agrochemical application (13.5%). At the close reference site, agrochemical application contributed 42.8%, followed by irrigation (22.7%), natural sources (17.4%), and mining activities (17.2%). At the far reference site, agrochemical application was predominant (40.6%), followed by irrigation (32.5%), and natural sources (26.9%). In comparison, at the mine site and the close reference site in sugarcane ecosystems, agrochemical application was predominant (50.1% and 57.4%, respectively), followed by mining activities (49.9% and 42.6%). At the far reference site, agrochemical application contributed 51.2%, followed by natural sources (48.8%). Therefore, the PMF model indicated that the optimal solution was four or three sources per site for rice paddies, but only two sources per site for sugarcane, suggesting that sources of metal/metalloid contamination were more complicated in rice paddy than in sugarcane agroecosystems.

Mixed-species bird flocks re-assemble interspecific associations across an elevational gradient.

Understanding how non-trophic social systems respond to environmental gradients is still a challenge in animal ecology, particularly in comparing changes in species composition to changes in interspecific interactions. Here, we combined long-term monitoring of mixed-species bird flocks, data on participating species' evolutionary history and traits, to test how elevation affected community assemblages and interspecific interactions in flock social networks. Elevation primarily affected flocks through reassembling interspecific associations rather than modifying community assemblages. Specifically, flock networks at higher elevations (compared to low elevations) had stronger interspecific associations (larger average weighted degree), network connectivity (enhanced network density) and fewer subnetworks. A phylogenetic and functional perspective revealed that associations between similar species weakened, whereas connections between dissimilar and/or random species were unchanged or strengthened with elevation. Likewise, network assortativity for the traits of vertical stratum and breeding period declined with elevation. The overall pattern is a change from modular networks in the lowlands, where species join flocks with other species that have matching traits, to a more open, random system at high elevations. Collectively, this rewiring of interspecific networks across elevational gradients imparts network stability and resiliency and makes mixed-species flocks less sensitive to local extinctions caused by harsh environments.

Increasing collaboration between China and India in the environmental sciences to foster global sustainability.

As the two largest countries by population, China and India have pervasive effects on the ecosphere. Because of their human population size and long international boundary, they share biodiversity and the threats to it, as well as crops, pests and diseases. We ranked the two countries on a variety of environmental challenges and solutions, illustrating quantitatively their environmental footprint and the parallels between them regarding the threats to their human populations and biodiversity. Yet we show that China and India continue to have few co-authorships in environmental publications, even as their major funding for scientific research has expanded. An agenda for collaboration between China and India can start with the shared Himalaya, linking the countries' scientists and institutions. A broader agenda can then be framed around environmental challenges that have regional patterns. Coordinated and collaborative research has the potential to improve the two countries' environmental performance, with implications for global sustainability.

Passerine bird reproduction does not decline in a highly-contaminated mercury mining district of China.

Mercury (Hg) is a neurotoxic element with severe effects on humans and wildlife. Widely distributed by atmospheric deposition, it can also be localized near point sources such as mines. Mercury has been shown to reduce the reproduction of bird populations in field observations in North America and Europe, but studies are needed in Asia, where the majority of emissions now occur. We investigated the reproduction of two passerines, Japanese Tit (Parus minor) and Russet Sparrow (Passer rutilans), in a large-scale Hg mining district, and a non-mining district, both in Guizhou, southwest China. Concentrations of Hg were elevated in the mining district (blood levels of 2.54 ± 2.21 [SD] and 0.71 ± 0.40 µg/g, in adult tits and sparrows, respectively). However, we saw no evidence of decreased breeding there: metrics such as egg volume, nestling weight, hatching and fledgling success, were all similar between the different districts across two breeding seasons. Nor were there correlations at the mining district between Hg levels of adults or juveniles, and hatching or fledgling success, or nestling weight. Nest success was high even in the mining district (tit, 64.0%; sparrow: 83.1%). This lack of reproductive decline may be related to lower blood levels in nestlings (means < 0.15 µg/g for both species). Concentrations of selenium (Se), and Se-to-Hg molar ratio, were also not correlated to breeding success. Although blood levels of 3.0 µg/g have been considered as a threshold of adverse effects in birds, even leading to severe effects, we detected no population-level reproductive effects, despite ~25% of the adult tits being above this level. Future work should investigate different locations in the mining district, different life-stages of the birds, and a wider variety of species. The hypothesis that bird populations can evolve resistance to Hg in contaminated areas should also be examined further.

Behavioural variables influence contact call rate more than characteristics of the vegetation in a group-living passerine species.

Although the contact calls of birds have been studied for their acoustic properties, limited research has investigated their repetitive nature. The rate of contact calls could be related to movement, with recruiting birds signalling their location, or it could help maintaining spacing between group mates, or give information about the environment where both signaller and receiver are located. If maintaining spacing, higher call rates would be expected in denser vegetation; alternatively, if birds gain information about predation risk from the cessation of contact calling, then open areas might elicit higher call rate. We studied how contact call rate in groups of Swinhoe's White-eyes (Zosterops simplex) was influenced by vegetation, collecting a total of 800 recordings. After statistically controlling for group size, the vegetation effect was weak and inconsistent. However, flying individuals produced a distinct flight call consisting of repeated notes similar to contact calls, and group-level contact call rate increased before flights, particularly when birds flew into the group. Therefore, we believe that contact call rate indicates information about individual or group movements, and could function as a continuous signal about the need for recruitment. We encourage further studies investigating how habitat, risk and audience influence contact call rate.

Endemicity and land-use type influence the abundance-range-size relationship of birds on a tropical island.

A single adverse environment event can threaten the survival of small-ranged species while random fluctuations in population size increase the extinction risk of less-abundant species. The abundance-range-size relationship (ARR) is usually positive, which means that smaller-ranged species are often of low abundance and might face both problems simultaneously. The ARR has been reported to be negative on tropical islands, perhaps allowing endemic species in such environments to remain extant. But there is a need to understand how endemism and land-use interact to shape ARR. Using 41 highly replicated transects along the full elevational gradient of Sri Lanka, we determined the following: (a) the direction of ARR, (b) if endemism affects ARR and (c) if land-use (rainforest, buffer and agriculture) changes ARR differently for endemics and non-endemics. Additionally, (d) we identified endemics that had both lower abundances and smaller range sizes, and ranked them from most threatened (specific to rainforests) to least threatened using a weighted-interaction nestedness estimator. (a) We found a positive relationship between species abundances and range size. This positive ARR was maintained among endemic and non-endemic species, across land-use types and at local and regional scales. (b) The ARR interacted with endemicity and land-use. Endemics with smaller range sizes had higher abundances than non-endemics, and particularly higher in rainforests compared to agriculture. In contrast, species with larger range sizes had similar abundances across endemicity and land-use categories. Many endemics with smaller range sizes are globally threatened; therefore, higher abundances may buffer them from extinction risks. (c) Nine (29%) endemics had both below average abundance and elevational range size. The nestedness estimator ranked the endemics Sri Lanka Whistling Thrush Myophonus blighi, Red-faced Malkoha Phaenicophaeus pyrrhocephalus, Sri Lanka Thrush Zoothera imbricata and White-faced Starling Sturnornis albofrontus as the four most vulnerable species to local extinction risk, which corresponds to their global extinction risk. We demonstrate that ARR can be positive on tropical islands, but it is influenced by endemism and land-use. Examining shifts in ARR is not only important to understand community dynamics but can also act as a tool to inform managers about species that require monitoring programmes.

Strictly protected areas are not necessarily more effective than areas in which multiple human uses are permitted.

The International Union for Conservation of Nature (IUCN) classifies protected areas into six categories, ranging from strict nature reserves to areas where multiple human uses are permitted. In the past, many researchers have questioned the effectiveness of multiple-use areas, fueling an unresolved debate regarding their conservation value. The literature so far has been inconclusive: although several studies have found that strictly protected areas are more effective, others have found the opposite, and yet others that the two types do not differ. To help resolve this debate, we reviewed the literature on protected areas and conducted our own analysis using > 19 000 terrestrial protected areas worldwide. We found that the differences between strictly protected areas and areas in which multiple human uses are permitted are often small and not statistically significant. Although the effectiveness of protected areas worldwide varies, other factors, besides their assigned IUCN category, are likely to be driving this pattern.